Hydrological niche segregation and species coexistence in arid zones

doi:10.7522/j.issn.1000-694X.2025.00364

Journal of Desert Research ›› 2026, Vol. 46 ›› Issue (1): 175-183.DOI: 10.7522/j.issn.1000-694X.2025.00364

Hydrological niche segregation and species coexistence in arid zones

Hai Zhou¹(), Wenzhi Zhao¹, Zhibin He¹, Lihui Tian², Huli Gu¹, Mingyan Fan¹^,²

^1.Linze Inland River Basin Research Station / State Key Laboratory of Ecological Safety and Sustainable Development in Arid Lands，Northwest Institute of Eco-Environment and Resources，Chinese Academy of Sciences，Lanzhou 730000，China
^2.State Key Laboratory of Plateau Ecology and Agriculture，Qinghai University，Xining 810016，China

Received:2025-11-23 Revised:2025-12-31 Online:2026-01-20 Published:2026-03-09

Abstract

Abstract:

In arid regions， water availability serves as a critical limiting factor governing the stability of plant communities and the functioning of ecosystems. To cope with drought stress， plants have developed diverse survival strategies through long-term evolutionary processes. Species diversity within plant communities often relies on niche differentiation driven by spatiotemporal heterogeneity in water availability， whereby coexisting species exhibit distinct growth and survival strategies along hydrological gradients. This paper systematically elucidates the core principles of the water niche hypothesis， emphasizing that hydrological niche segregatio enables plants to utilize limited water resources differentially through variations in traits such as root distribution， phenological responses， and stomatal regulation. Spatially， water niche differentiation manifests as vertical stratification of root systems driven by soil moisture heterogeneity； temporally， it leads to asynchronous species responses via the "storage effect." Furthermore， interspecific variation in stomatal traits provides a physiological foundation for this differentiation. In summary， hydrological niche segregatio reduces direct interspecific competition for water resources， thereby functioning as a key mechanism promoting stable coexistence among plant species， maintaining community diversity， and ensuring ecosystem stability in arid regions. Advances in techniques such as stable isotope analysis offer robust tools for quantifying niche differentiation. A deeper understanding of this mechanism holds significant theoretical and practical implications for vegetation restoration and ecosystem management in arid zones.

Key words: desert vegetation, hydrological niche segregation, species coexistence, spatiotemporal heterogeneity, stomatal regulation

CLC Number:

Hai Zhou, Wenzhi Zhao, Zhibin He, Lihui Tian, Huli Gu, Mingyan Fan. Hydrological niche segregation and species coexistence in arid zones[J]. Journal of Desert Research, 2026, 46(1): 175-183.

References 103

[1]	Haase P， Pugnaire F I， Clark S C，et al.Environmental control of canopy dynamics and photosynthetic rate in the evergreen tussock grass Stipa tenacissima ［J］.Plant Ecology，1999，145（2）：327-339.
[2]	Dube O P， Pickup G.Effects of rainfall variability and communal and semi-commercial grazing on land cover insouthern African ranglands［J］.Climate Res，2001，17（2）：195-208.
[3]	赵文智，郑颖，张格非.绿洲边缘人工固沙植被自组织过程［J］.中国沙漠，2018，38（1）：1-7.
[4]	Yao S， Hu W， Ji M，et al.Distribution，species richness，and relative importance of different plant life forms across drylands in China［J］.Plant Diversity，2025，47（2）：273-281.
[5]	Bernacchi C J， VanLoocke A.Terrestrial ecosystems in a changing environment：a dominant role for water［J］.Annual Review of Plant Biology，2015，66：599-622.
[6]	Lambers H， Oliveira R S.Plant water relations［M］//Lambers H，Oliveira R S.Plant Physiological Ecology.Cham，Germany：Springer International Pblishing，2019：187-263.
[7]	Dawson T E， Ehleringer J R.Streamside trees that do not use stream water［J］.Nature，1993，150（6316）：335-337.
[8]	Asbjornsen H， Goldsmith G R， Alvarado-Barrientos M S，et al.Ecohydrological advances and applications in plant-water relations research：a review［J］.Journal of Plant Ecology，2011，4（1）：3-22.
[9]	Silvertown J， Araya Y， Gowing D.Hydrological niches in terrestrial plant communities：a review［J］.Journal of Ecology，2015，103（1）：93-108.
[10]	Jobbágy E G， Nosetto M D， Villagra P E，et al.Water subsidies from mountains to deserts：their role in sustaining groundwater‐fed oases in a sandy landscape［J］.Ecological Applications，2011，21（3）：678-694.
[11]	García‐Baquero G， Silvertown J， Gowin D J，et al.Dissecting the hydrological niche：soil moisture，space and lifespan［J］.Journal of Vegetation Science，2016，27（2）：219-226.
[12]	Schwinning S， Ehleringer J R.Water use trade-offs and optimal adaptations to pulse-driven arid ecosystems［J］.Journal of Ecology，2001，89：464-480.
[13]	West A G， Dawson T E， February E C，et al.Diverse functional responses to drought in a Mediterranean-type shrubland in South Africa［J］.New Phytologist，2012，195：396-407.
[14]	Letten A D， Keith D A， Tozer M G，et al.Fine‐scale hydrological niche differentiation through the lens of multispecies cooccurrence models［J］.Journal of Ecology，2015，103（5）：1264-1275.
[15]	Navas M L， Violle C.Plant traits related to competition：how do they shape the functional diversity of communities？［J］.Community Ecology，2009，10（1）：131-137.
[16]	Laliberté E， Zemunik G， Turner B L.Environmental filtering explains variation in plant diversity along resource gradients［J］.Science，2014，345（6204）：1602-1605.
[17]	Huxman T E， Venable D L.Understanding past，contemporary，and future dynamicsof plants，populations，and communities using Sonoran Desert winter annuals［J］.American Journal of Botany，2013，100：1369-1380.
[18]	Siepielski A M， McPeek M A.On the evidence for species coexistence：a critique of the coexistence program［J］.Ecology，2010，91（11）：3153-3164.
[19]	储诚进，王酉石，刘宇，等.物种共存理论研究进展［J］.生物多样性，2017，25：345-354.
[20]	Chesson P.Mechanisms of maintenance of species diversity［J］.Annual Review of Ecology and Systematics，2000，31：343-366.
[21]	Letten A D， Ke P J， Fukami T.Linking modern coexistence theory and contemporary niche theory［J］.Ecological Monographs，2017，87：161-177.
[22]	Chesson P， Gebauer R L， Schwinning S，et al.Resource pulses，species interactions，and diversity maintenance in arid and semi-arid environments［J］.Oecologia，2004，141（2）：236-253.
[23]	Orians G H， Solbrig O T.A cost-income model of leaves and roots with special reference to arid and semiarid areas［J］.The American Naturalist，1977，111：677-690.
[24]	Peñuelas J， Terradas J， Lloret F.Solving the conundrum of plant species coexistence：water in space and time matters most［J］.New Phytologist，2011，189：5-8.
[25]	Dawson T E.Spatial and physiological overlap of three co-occurring alpine willows［J］.Functional Ecology，1990，4：13-25.
[26]	Silvertown J， Dodd M E， Gowing D J G，et al.Hydrologically defined niches reveal a basis for species richness in plant communities［J］.Nature，1999，400：61-63.
[27]	Weissert H.Global change：deciphering methane's fingerprint［J］.Nature，2000，406（6794）：356-357.
[28]	Gibson J J， Reid R.Stable isotope fingerprint of open-water evaporation losses and effective drainage area fluctuations in a subarctic shield watershed［J］.Journal of Hydrology，2010，381（1）：142-150.
[29]	Craig H， Gordon L I.Deuterium and oxygen 18 variations in the ocean and the marine atmospher［C］.Symposium on Marine Geochemistry.1964.
[30]	Dawson T E， Mambelli S， Plamboeck A H，et al.Stable isotopes in plant ecology［J］.Annual Review of Ecology and Systematics，2002，33：507-559.
[31]	O'Keefe K， Nippert J B， McCulloh K A.Plant water uptake along a diversity gradient provides evidence for complementarity in hydrological niches［J］.Oikos，2019，128：1748-1760.
[32]	Moreno-Gutiérrez C， Dawson T E， Nicolas E，et al.Isotopes reveal contrasting water use strategies among coexisting plant species in a Mediterranean ecosystem［J］.New Phytologist，2012，196：489-496.
[33]	Volkmann T， Kühnhammer K， Herbstritt B，et al.A method for in situ monitoring of the isotope composition of tree xylem water using laser spectroscopy［J］.Plant，Cell Environment，2016，39（9）：2055-2063.
[34]	Beyer M， Dubbert M.Water worlds and how to investigate them：a review and future perspective on in situ measurements of water stable isotopes in soils and plants［J］.Hydrology and Earth System Sciences Discussions，2019：1812-2108.
[35]	Kühnhammer K， Dahlmann A， Iraheta A，et al.Continuous in situ measurements of water stable isotopes in soils，tree trunk and root xylem：field approval［J］.Rapid Communications in Mass Spectrometry，2022，36（5）：e9232.
[36]	Holloway-Phillips M， Cernusak LA， Barbour M，et al.Leaf vein fraction influences the Péclet effect and ¹⁸O enrichment in leaf water［J］.Plant，Cell and Environment，2016，39：2414-2427.
[37]	Dubbert M， Cuntz M， Piayda A，et al.Oxygen isotope signatures of transpired water vapor：the role of isotopic non‐steady‐state transpiration under natural conditions［J］.New Phytologist，2014，203（4）：1242-1252.
[38]	Araya Y N， Silvertown J， Gowing D J，et al.A fundamental，eco-hydrological basis for niche segregation in plant communities［J］.New Phytologist，2011，189：253-258.
[39]	Johnson R H.Determinate evolution in the color-pattern of the lady-beetles［J］.Washington Carnegie Inst，1910.
[40]	Grinnell J.The niche relationships of the California thrasher［J］.AUK，1917，34：427-433.
[41]	Leibold M A.Similarity and local co-existence of species in regional biotas［J］.Evolutionary Ecology，1998，12：95-110.
[42]	Kylafis G， Loreau M.Niche construction in the light of niche theory［J］.Ecology Letters，2011，14（2）：82-90.
[43]	Yamamichi M， Kyogoku D， Iritani R，et al.Intraspecific adaptation load：a mechanism for species coexistence［J］.Trends in Ecology Evolution，2020，35（10）：897-907.
[44]	Ivanov V Y， Hutyra L R， Wofsy S C，et al.Root niche separation can explain avoidance of seasonaldrought stress and vulnerability of overstory trees to extended drought in a mature Amazonian forest［J］.Water Resources Research，2012，48：W12507.
[45]	Huxman T E， Smith M D， Fay P A，et al.Convergence across biomes to a common rain-use efficiency［J］.Nature，2004，429（6992）：651-654.
[46]	Levine J M， HilleRisLambers J.The importance of niches for the maintenance of species diversity［J］.Nature，2009，461：254-258.
[47]	Brum M， Vadeboncoeur M A， Ivanov V，et al.Hydrological niche segregation defines forest structure and drought tolerance strategies in a seasonal Amazon forest［J］.Journal of Ecology，2018，107（1）：318-333.
[48]	Silvertown J.Plant coexistence and the niche［J］.Trends in Ecology and Evolution，2004，19：605-611.
[49]	Stein A， Gerstner K， Kreft H.Environmental heterogeneity as a universal driver of species richness across taxa，biomes and spatial scales［J］.Ecology Letters，2014，17：866-880.
[50]	Adler P B， Fajardo A， Kleinhesselink A R，et al.Traitbased tests of coexistence mechanisms［J］.Ecology Letters，2013，16：1294-1306.
[51]	Kraft N J B， Adler P B， Godoy O，et al.Community assembly，coexistence，and the environmental filtering metaphor［J］.Functional Ecology，2014，29：592-599.
[52]	Grubb P J.The maintenance of species-richness in plant communities：the importance of the regeneration niche［J］.Biological Reviews，1977，52：107-145.
[53]	Amarasekare P.Competitive coexistence in spatially structured environments：a synthesis［J］.Ecology Letters，2003，6：1109-1122.
[54]	Gómez‐Plaza A， Alvarez-Rogel J， Albaladejo J，et al.Spatial patterns and temporal stability of soil moisture across a range of scales in a semi-arid environment［J］.Hydrological Processes，2000，14（7）：1261-1277.
[55]	Wilson D J， Western A W， Grayson R B.Identifying and quantifying sources of variability in temporal and spatial soil moisture observations［J］.Water Resources Research，2004，40（2）：191-201.
[56]	李新荣，张志山，谭会娟，等.我国北方风沙危害区生态重建与恢复：腾格里沙漠土壤水分与植被承载力的探讨［J］.中国科学：生命科学，2014，44：257-266.
[57]	Shao M A， Wang Y， Xia Y，et al.Soil drought and water carrying capacity for vegetation in the critical zone of the Loess Plateau：a review［J］.Vadose Zone Journal，2018，17（1）：1-8.
[58]	Flanagan L B， Ehleringer J R， Marshall J D.Differential uptake of summer precipitation among co-occurring trees and shrubs in a pinyon-junipe woodland［J］.Plant Cell and Environment，1992，15：831-836.
[59]	Xu H， Li Y， Xu G Q，et al.Ecophysiological response and morphological adjustment of two Central Asian desert shrubs towards variation in summer precipitation［J］.Plant，Cell and Environment，2007，30：399-409.
[60]	Smucher A J M， Ritchie J T.Plant physiological responses to the soil［J］.International Crop Science，1993：747-748.
[61]	Jean-François Barczi， Hervé R， Sébastien G，et al.DigR：a generic model and its open source simulation software to mimic three-dimensional root-system architecture diversity.［J］Annals of Botany，2018，121（5）：1089-1104.
[62]	Hutchings M J， John E A， Wijesinghe D K.Toward understanding the consequences of soilheterogeneity for plant populations and communities［J］.Ecology，2003，84：2322-2334.
[63]	Postma J A， Hecht V L， Hikosaka K，et al.Dividing the pie：a quantitative review on plant density responses［J］.Plant，Cell and Environment，2021，44（4）：1072-1094.
[64]	de Kroon H， Hendriks M， van Ruijven J，et al.Root responses to nutrients and soil biota：drivers of species coexistence and ecosystem productivity［J］.Journal of Ecology，2012，100（1）：6-15.
[65]	Wang P， Shu M， Mou P.Fine root responses to temporal nutrient heterogeneity and competition in seedlings of two tree species with different rooting strategies［J］.Ecology and Evolution，2018，8（6）：3367-3375.
[66]	Rajaniemi T K， Allison V J， Goldberg D E.Root competition can cause a decline in diversity with increased productivity［J］.Journal of Ecology，2003，91：407-416.
[67]	Mahaut L， Fort F， Violle C.Multiple facets of diversity effects on plant productivity：species richness，functional diversity，species identity and intraspecific competition［J］.Functional Ecology，2020，34（1）：287-298.
[68]	Angert A L， Huxman T E， Chesson P，et al.Functional tradeoffs determine species coexistence via the storage effect［J］.Proceedings of the National Academy of Sciences of the United States of America，2009，106：11641-11645.
[69]	Usinowicz J， Chang-Yang C H， Chen Y Y，et al.Temporal coexistence mechanisms contribute to the latitudinal gradient in forest diversity［J］.Nature，2017，550：105-108.
[70]	潘元琪，杜彦君，陈文德，等.植物物候是否能解释物种共存：以浙江古田山亚热带常绿阔叶林为例［J］.中国科学：生命科学，2020，50：362-372.
[71]	王阿晴，朱雅娟，马媛，等.戈壁沙冬青（Ammopiptanthus mongolicus）群落水分来源的年际动态［J］.中国沙漠，2025，45（1）：85-93.
[72]	Schwinning S， Sala O E.Hierarchy of responses to resource pulses in and semiarid ecosystem［J］.Oecologia，2004，141：211-220.
[73]	Dixit A， Kolb T， Burney O.Trade-off between growth rate and water use efficiency in southwestern ponderosa pine provenances［J］.Forest Ecology and Management，2022，515：120239.
[74]	Verhulst J， Montana C， Mandujano M C，et al.Demographic mechanisms in the coexistence of two closely related perennials in a fluctuating environment［J］.Oecologia，2008，156：95-105.
[75]	Overpeck J T， Udall B.Climate change and the aridification of North America［J］.Proceedings of the National Academy of Sciences，2020，117（22）：11856-11858.
[76]	Terradas J， Penuelas J， Lloret F.The fluctuation niche in plants［J］.International Journal of Ecology，2009：1-5.
[77]	王海仙，张勇勇，康文蓉，等.干旱半干旱区典型植被类型蒸散比及土壤水分阈值［J］.中国沙漠，2025，45（6）：289-299.
[78]	Chen X Y.Seasonal patterns of fine-root productivity and turnover in a tropical savanna of northern Australia［J］.Journal of Tropical Ecology，2004，20：221-224.
[79]	Fort F， Volaire F， Guilioni L，et al.Root traits are related to plant water-use amongrangeland Mediterranean species［J］.Functional Ecology，2017，31：1700-1709.
[80]	Li Z， Lamb E G， Piper C L，et al.Plant belowground diversity and species segregation by depth in a semi-arid grassland［J］.Écoscience，2018，25（1）：1-7.
[81]	Sack L， Buckley T N.The developmental basis of stomatal density and flux［J］.Plant Physiology，2016，171（4）：2358-2363.
[82]	Liu C， Sack L， Li Y，et al.Relationships of stomatal morphology to the environment across plant communities［J］.Nature Communications，2023，14（1）：6629.
[83]	Xie J， Wang Z， Li Y.Stomatal opening ratio mediates trait coordinating network adaptation to environmental gradients［J］.New Phytologist，2022，235（3）：907-922.
[84]	Wright I J， Reich P B， Westoby M，et al.The worldwide leaf economics spectrum［J］.Nature，2004，428：821-827.
[85]	Lawson T， Vialet-Chabrand S.Speedy stomata，photosynthesis and plant water use efficiency［J］.New Phytologist，2019，221（1）：93-98.
[86]	Pan S， Wang X， Yan Z，et al.Leaf stomatal configuration and photosynthetic traits jointly affect leaf water use efficiency in forests along climate gradients［J］.New Phytologist，2024，244（4）：1250-1262.
[87]	Seibt U， Rajabi A， Griffiths H，et al.Carbon isotopes and water use efficiency：sense and sensitivity［J］.Oecologia，2008，155：441-454.
[88]	Tardieu F， Simonneau T.Variability among species of stomatal control under fluctuating soil water status and evaporative demand：modelling isohydric and anisohydric behaviours［J］.Journal of Experimental Botany，1998，49：419-432.
[89]	West A G， Hultine K R， Jackson T L，et al.Differential summer water use by Pinus edulis and Juniperus osteosperma reflects contrasting hydraulic characteristics［J］.Tree Physiology，2007，27：1711-1720.
[90]	McDowell N， Pockman W T， Allen C D，et al.Mechanisms of plant survival and mortality during drought：why do some plants survive while others succumb to drought？［J］.New Phytologist，2008，178：719-739.
[91]	Klein T， Cohen S， Yakir D.Hydraulic adaptations underlying drought resistance of Pinus halepensis ［J］.Tree Physiology，2011，31：637-648.
[92]	Jackson R B， Jobbagy E G， Avissar R，et al.Trading water for carbon with biological carbon sequestration［J］.Science，2005，310：1944-1947.
[93]	Farely K A， Jobbagy E G， Jackson R B.Effects of afforestation on water yield：a global synthesis with implications for policy［J］.Global Change Biol，2005，11：1565-1576.
[94]	Grunzweig J M， Gelfand I， Yakir D.Biogeochemical factors contributing to enhanced carbon storage following afforestation of a semi-arid shrubland［J］.Biogeosciences，2007，4：891-904.
[95]	Baldocchi D D， Ma S， Rambal S，et al.On the differential advantages of evergreenness and deciduousness in Mediterranean oak woodlands：a flux perspective［J］.Ecological Applications，2010，20：1583-1597.
[96]	Schultz H R.Differences in hydraulic architecture account for near-isohydric and anisohydric behavior of two field-grown Vitis vinifera cultivars during drought［J］.Plant，Cell & Environment，2003，26：1393-1405.
[97]	Mereu S， Salvatori E， Fusaro L，et al.A whole plant approach to evaluate the water use of Mediterranean maquis species in a coastal dune ecosystem［J］.Biogeosci.Disscuss，2009，6：1713-1746.
[98]	Van der Molen M K， Dolmana A J， Ciais P，et al.Drought and ecosystem carbon cycling［J］.Agricultural and Forest Meteorology，2011，151：765-773.
[99]	Keddy P.Plants and Vegetation：Origins，Processes，Consequences［M］.Cambridge，UK：Cambridge University Press，2007.
[100]	Zhao Y， Wang L， Chun K P，et al.Dynamic hydrological niche segregation：How plants compete for water in a semi-arid ecosystem［J］.Journal of Hydrology，2024，630：130677.
[101]	Gu H， Zhou H， Chen G，et al.Isotopic evidence of hydrological niche segregation and species coexisting mechanism of desert shrubs Reaumuria songarica and Nitraria sphaerocarpa ［J］.Journal of Hydrology，2025：133803.
[102]	Fan M， Zhou H， Tian L，et al.Hydrological niche separation between two coexisting shrubs in an extremely arid region［J］.Journal of Hydrology：Regional Studies，2025，57：102170.
[103]	Cai L， Xiong K， Li Y，et al.Coexisting plants restored in karst desertification areas cope with drought by changing water uptake patterns and improving water use efficiency［J］.Journal of Hydrology，2025，654：132813.

Hydrological niche segregation and species coexistence in arid zones

RichHTML

PDF

Knowledge

Abstract

Cite this article

share this article

References 103

Related Articles 11

Recommended Articles

Metrics

Comments

[1]	Jun Lei, Xiaohu Yang, Hongmei Liu, Yuhong Zhao, Juping Fan, Caixia Guo. The characteristics of desert vegetation biomass and its influencing factors in the middle reaches of the Heihe River [J]. Journal of Desert Research, 2021, 41(1): 203-208.
[2]	Mengyu Hao, Longjun Qin, Peng Mao, Jiechunyi Luo, Wenli Zhao, Guoyu Qiu. Unmanned aerial vehicle （UAV） based methodology for spatial distribution pattern analysis of desert vegetation [J]. Journal of Desert Research, 2020, 40(6): 169-179.
[3]	Lei Wu, Changbin Li, Liuming Wang, Xuhong Xie, Yuan Zhang, Jianmei Wei. Division and application of desert-oasis system in arid Northwest China based on ESA-LUC and MODIS-NDVI [J]. Journal of Desert Research, 2020, 40(6): 139-150.
[4]	Li Yike, Zhao Chengyi, Yang Ruihong. Intraspecific Competition of Haloxylon ammodendron in the Southern Margin of Junggar Basin [J]. JOURNAL OF DESERT RESEARCH, 2016, 36(2): 335-341.
[5]	Zhong Zebing, Zhou Guoying, Yang Lucun, Liu Hechun, Song Wenzhu. The Biomass Allocation Patterns of Desert Shrub Vegetation in the Qaidam Basin, Qinghai, China [J]. JOURNAL OF DESERT RESEARCH, 2014, 34(4): 1042-1048.
[6]	ZHAO Jin;CHEN Xi;Guli Japper;MA Zhong-guo;CHANG Cun;. Spectral Discrimination of Desert Vegetation in the Tarim River Basin [J]. JOURNAL OF DESERT RESEARCH, 2009, 29(2): 270-278.
[7]	ZHANG Kai;SI Jian-hua;WANG Run-yuan;WANG Xiao-ping;HAN Hai-tao;GUO Ni. Impact of Climate Change on Desert Vegetation in Alxa Region [J]. JOURNAL OF DESERT RESEARCH, 2008, 28(5): 879-885.
[8]	ZHANG Jin-chun;ZHAO Ming;LIAO Kong-tai;XU Yan-shuang;ZHANG Ying-chang. Experimental Research on Desert Vegetation Restoration by Drip Irrigation at Minqin Oasis Edge [J]. JOURNAL OF DESERT RESEARCH, 2007, 27(1): 94-98.
[9]	WANG Ji-he;MA Quan-lin;LIU Hu-jun;YANG Zi-hui;ZHANG De-kui. Effect of Wind-breaking and Sand-fixing of Vegetation in Progressive Succession on Desertification Land in Arid Area [J]. JOURNAL OF DESERT RESEARCH, 2006, 26(6): 908-909.
[10]	XIAO Sheng-chun;XIAO Hong-lang;SONG Yao-xuan;DUAN Zheng-hu;LU Ming-feng. Dendrochronology Study on Response of Reaumurta soongorica to Water-Heat Variation [J]. JOURNAL OF DESERT RESEARCH, 2006, 26(4): 548-552.
[11]	YANG Zi hui. Research on Desert Vegetation Changes for 40 Years at Shajingzi Area in Minqin [J]. JOURNAL OF DESERT RESEARCH, 1999, 19(4): 395-398.